Fluctuating selection

Last updated
Alleles A and B fluctuate in adaptive value over time Fluctuating selection.png
Alleles A and B fluctuate in adaptive value over time

Fluctuating selection is a mode of natural selection characterized by the fluctuation of the direction of selection on a given phenotype over a relatively brief period of evolutionary time. For example, a species of plant may come in two varieties: one which prefers wetter soil and one which prefers dryer soil. During a period of wet years, the wet variety will be more fit and produce more offspring, and thereby increase the frequency of wet-preferring plants. If this wet period is followed by drought, the dry variety will be selected for and its numbers will increase. As periods of dryness and wetness fluctuate, so too does selection on dry-preferring and wet-preferring plants. Fluctuating selection is also manifest at the genic level. Consider two alleles, A and B, which are found at the same locus. Fluctuating selection dynamics are at play when selection favors A at time t0, B at t1 and A again at t2.

Contents

Fluctuating selection has been characterized by several mathematical models. [1] [2] [3] Under some circumstances, fluctuating selection may lead to a balanced polymorphism. [4] When two species exert selection on one another, e.g. a host and its parasite, this can lead to fluctuating selection dynamics. [5] [6] [7]

Red Queen dynamics and the maintenance of sex

The Red Queen hypothesis describes coevolutionary 'arms races' between antagonistic species (predators and prey, parasites and hosts, competitors with overlapping niches), emphasizing competition between species and populations rather than within them. Under Red Queen dynamics, a species must adapt to shifting selection pressures of the ever-changing biota which constitute its environment or face extinction. Experiments in Red Queen environments on real and simulated populations have offered strong support for the maintenance of sexual reproduction despite the two-fold cost of sex. [8] [9]

Fluctuating selection may also play an important role in host-parasite coevolutionary relationships, specifically in the maintenance of sex. It has been shown that coevolutionary arms race dynamics between host and parasite give way to fluctuating selection dynamics in a minimal environment. [6] Fluctuating selection in Red Queen environments has been suggested as an explanation for the persistence of sex:

The essence of sex in our theory is that it stores genes that are currently bad but have promise for reuse. It continually tries them in combination, waiting for the time when the focus of disadvantage has moved elsewhere. When this has happened, the genotypes carrying such genes spread by successful reproduction, becoming simultaneously stores for other bad genes and thus onward in continuous succession. [8]

In this conception of sex, the population is a storehouse of variation and sex is a mechanism for distributing old, minority variants once they become useful. This theory depends on fluctuating selection, as fluctuating selection dynamics make adaptive previously maladaptive variants due to ecological shifts.

See also

Related Research Articles

<span class="mw-page-title-main">Evolution</span> Change in the heritable characteristics of biological populations

Evolution is the change in the heritable characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, resulting in certain characteristics becoming more or less common within a population over successive generations. The process of evolution has given rise to biodiversity at every level of biological organisation.

<span class="mw-page-title-main">Sexual selection</span> Mode of natural selection involving the choosing of and competition for mates

Sexual selection is a mechanism of evolution in which members of one biological sex choose mates of the other sex to mate with, and compete with members of the same sex for access to members of the opposite sex. These two forms of selection mean that some individuals have greater reproductive success than others within a population, for example because they are more attractive or prefer more attractive partners to produce offspring. Successful males benefit from frequent mating and monopolizing access to one or more fertile females. Females can maximise the return on the energy they invest in reproduction by selecting and mating with the best males.

<span class="mw-page-title-main">Parasitism</span> Relationship between species where one organism lives on or in another organism, causing it harm

Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson characterised parasites as "predators that eat prey in units of less than one". Parasites include single-celled protozoans such as the agents of malaria, sleeping sickness, and amoebic dysentery; animals such as hookworms, lice, mosquitoes, and vampire bats; fungi such as honey fungus and the agents of ringworm; and plants such as mistletoe, dodder, and the broomrapes.

Selfish genetic elements are genetic segments that can enhance their own transmission at the expense of other genes in the genome, even if this has no positive or a net negative effect on organismal fitness. Genomes have traditionally been viewed as cohesive units, with genes acting together to improve the fitness of the organism.

<span class="mw-page-title-main">Coevolution</span> Two or more species influencing each others evolution

In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well as gene-culture coevolution.

<span class="mw-page-title-main">Muller's ratchet</span> Accumulation of harmful mutations

In evolutionary genetics, Muller's ratchet is a process which, in the absence of recombination, results in an accumulation of irreversible deleterious mutations. This happens because in the absence of recombination, and assuming reverse mutations are rare, offspring bear at least as much mutational load as their parents. Muller proposed this mechanism as one reason why sexual reproduction may be favored over asexual reproduction, as sexual organisms benefit from recombination and consequent elimination of deleterious mutations. The negative effect of accumulating irreversible deleterious mutations may not be prevalent in organisms which, while they reproduce asexually, also undergo other forms of recombination. This effect has also been observed in those regions of the genomes of sexual organisms that do not undergo recombination.

<span class="mw-page-title-main">Genetic diversity</span> Total number of genetic characteristics in a species

Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species. It ranges widely, from the number of species to differences within species, and can be correlated to the span of survival for a species. It is distinguished from genetic variability, which describes the tendency of genetic characteristics to vary.

<span class="mw-page-title-main">Polymorphism (biology)</span> Occurrence of two or more clearly different morphs or forms in the population of a species

In biology, polymorphism is the occurrence of two or more clearly different morphs or forms, also referred to as alternative phenotypes, in the population of a species. To be classified as such, morphs must occupy the same habitat at the same time and belong to a panmictic population.

<span class="mw-page-title-main">Directional selection</span> Type of genetic selection favoring one extreme phenotype

In population genetics, directional selection is a type of natural selection in which one extreme phenotype is favored over both the other extreme and moderate phenotypes. This genetic selection causes the allele frequency to shift toward the chosen extreme over time as allele ratios change from generation to generation. The advantageous extreme allele will increase as a consequence of survival and reproduction differences among the different present phenotypes in the population. The allele fluctuations as a result of directional selection can be independent of the dominance of the allele, and in some cases if the allele is recessive, it can eventually become fixed in the population.

<span class="mw-page-title-main">Evolution of sexual reproduction</span>

Evolution of sexual reproduction describes how sexually reproducing animals, plants, fungi and protists could have evolved from a common ancestor that was a single-celled eukaryotic species. Sexual reproduction is widespread in eukaryotes, though a few eukaryotic species have secondarily lost the ability to reproduce sexually, such as Bdelloidea, and some plants and animals routinely reproduce asexually without entirely having lost sex. The evolution of sexual reproduction contains two related yet distinct themes: its origin and its maintenance. Bacteria and Archaea (prokaryotes) have processes that can transfer DNA from one cell to another, but it is unclear if these processes are evolutionarily related to sexual reproduction in Eukaryotes. In eukaryotes, true sexual reproduction by meiosis and cell fusion is thought to have arisen in the last eukaryotic common ancestor, possibly via several processes of varying success, and then to have persisted.

Evolutionary game theory (EGT) is the application of game theory to evolving populations in biology. It defines a framework of contests, strategies, and analytics into which Darwinian competition can be modelled. It originated in 1973 with John Maynard Smith and George R. Price's formalisation of contests, analysed as strategies, and the mathematical criteria that can be used to predict the results of competing strategies.

<span class="mw-page-title-main">Viviparous lizard</span> Species of lizard

The viviparous lizard, or common lizard, is a Eurasian lizard. It lives farther north than any other species of non-marine reptile, and is named for the fact that it is viviparous, meaning it gives birth to live young. Both "Zootoca" and "vivipara" mean "live birth", in (Latinized) Greek and Latin respectively. It was called Lacerta vivipara until the genus Lacerta was split into nine genera in 2007 by Arnold, Arribas & Carranza.

<span class="mw-page-title-main">Mate choice</span> Mechanism for evolution

Mate choice is one of the primary mechanisms under which evolution can occur. It is characterized by a "selective response by animals to particular stimuli" which can be observed as behavior. In other words, before an animal engages with a potential mate, they first evaluate various aspects of that mate which are indicative of quality—such as the resources or phenotypes they have—and evaluate whether or not those particular trait(s) are somehow beneficial to them. The evaluation will then incur a response of some sort.

The Red Queen's hypothesis is a hypothesis in evolutionary biology proposed in 1973, that species must constantly adapt, evolve, and proliferate in order to survive while pitted against ever-evolving opposing species. The hypothesis was intended to explain the constant (age-independent) extinction probability as observed in the paleontological record caused by co-evolution between competing species; however, it has also been suggested that the Red Queen hypothesis explains the advantage of sexual reproduction at the level of individuals, and the positive correlation between speciation and extinction rates in most higher taxa.

Host–parasite coevolution is a special case of coevolution, where a host and a parasite continually adapt to each other. This can create an evolutionary arms race between them. A more benign possibility is of an evolutionary trade-off between transmission and virulence in the parasite, as if it kills its host too quickly, the parasite will not be able to reproduce either. Another theory, the Red Queen hypothesis, proposes that since both host and parasite have to keep on evolving to keep up with each other, and since sexual reproduction continually creates new combinations of genes, parasitism favours sexual reproduction in the host.

Ecoimmunology or Ecological Immunology is the study of the causes and consequences of variation in immunity. The field of ecoimmunology seeks to give an ultimate perspective for proximate mechanisms of immunology. This approach places immunology in evolutionary and ecological contexts across all levels of biological organization.

Interlocus sexual conflict is a type of sexual conflict that occurs through the interaction of a set of antagonistic alleles at two or more different loci, or the location of a gene on a chromosome, in males and females, resulting in the deviation of either or both sexes from the fitness optima for the traits. A co-evolutionary arms race is established between the sexes in which either sex evolves a set of antagonistic adaptations that is detrimental to the fitness of the other sex. The potential for reproductive success in one organism is strengthened while the fitness of the opposite sex is weakened. Interlocus sexual conflict can arise due to aspects of male–female interactions such as mating frequency, fertilization, relative parental effort, female remating behavior, and female reproductive rate.

<span class="mw-page-title-main">Major histocompatibility complex and sexual selection</span> Adaptive immune gene selection

Major histocompatibility complex (MHC) genes code for cell surface proteins that facilitate an organism's immune response to pathogens as well as its ability to avoid attacking its own cells. These genes have maintained an unusually high level of allelic diversity throughout time and throughout different populations. This means that for each MHC gene, many alleles consistently exist within the population, and many individuals are heterozygous at MHC loci.

<span class="mw-page-title-main">Evolving digital ecological network</span>

Evolving digital ecological networks are webs of interacting, self-replicating, and evolving computer programs that experience the same major ecological interactions as biological organisms. Despite being computational, these programs evolve quickly in an open-ended way, and starting from only one or two ancestral organisms, the formation of ecological networks can be observed in real-time by tracking interactions between the constantly evolving organism phenotypes. These phenotypes may be defined by combinations of logical computations that digital organisms perform and by expressed behaviors that have evolved. The types and outcomes of interactions between phenotypes are determined by task overlap for logic-defined phenotypes and by responses to encounters in the case of behavioral phenotypes. Biologists use these evolving networks to study active and fundamental topics within evolutionary ecology.

Disease ecology is a sub-discipline of ecology concerned with the mechanisms, patterns, and effects of host-pathogen interactions, particularly those of infectious diseases. For example, it examines how parasites spread through and influence wildlife populations and communities. By studying the flow of diseases within the natural environment, scientists seek to better understand how changes within our environment can shape how pathogens, and other diseases, travel. Therefore, diseases ecology seeks to understand the links between ecological interactions and disease evolution. New emerging and re-emerging infectious diseases are increasing at unprecedented rates which can have lasting impacts on public health, ecosystem health, and biodiversity.

References

  1. Haldane, J. B. S.; Jayakar, S. D. (1963). "Polymorphism due to selection of varying direction". Journal of Genetics. 58 (2): 237–242. doi:10.1007/BF02986143. ISSN   0022-1333. S2CID   26313306.
  2. Karlin, S.; Lieberman, Uri (1974). "Random temporal variation in selection intensities: Case of large population size". Theoretical Population Biology. 6 (3): 355–382. Bibcode:1974TPBio...6..355K. doi:10.1016/0040-5809(74)90016-1. ISSN   0040-5809. PMID   4460262.
  3. Bell, G. (2010). "Fluctuating selection: the perpetual renewal of adaptation in variable environments". Philosophical Transactions of the Royal Society of London B: Biological Sciences. 365 (1537): 87–97. doi:10.1098/rstb.2009.0150. ISSN   0962-8436. PMC   2842698 . PMID   20008388.
  4. Bertram, Jason; Masel, Joanna (20 March 2019). "Different mechanisms drive the maintenance of polymorphism at loci subject to strong versus weak fluctuating selection". Evolution. 73 (5): 883–896. doi:10.1111/evo.13719. hdl: 10150/632441 . PMID   30883731. S2CID   83461372.
  5. Hairson, N. G.; Dillon, T. A. (1990). "Fluctuating Selection and Response in a Population of Freshwater Copepods". Evolution. 44 (7): 1796–1805. doi:10.2307/2409508. ISSN   0014-3820. JSTOR   2409508. PMID   28567814.
  6. 1 2 Hall, A. R.; Scanlan, P. D.; Morgan, A. D.; Buckling, A. (2011). "Host-parasite coevolutionary arms races give way to fluctuating selection". Ecology Letters. 14 (7): 635–642. Bibcode:2011EcolL..14..635H. doi:10.1111/j.1461-0248.2011.01624.x. ISSN   1461-023X. PMID   21521436.
  7. Sielpielski, A. M.; DiBattista, J. D.; Evans, J. A.; Carlson, S. M. (2011). "Differences in the temporal dynamics of phenotypic selection among fitness components in the wild". Proceedings of the Royal Society of London B: Biological Sciences. 278 (1711): 1572–1580. doi:10.1098/rspb.2010.1973. ISSN   0962-8452. PMC   3081754 . PMID   21047862.
  8. 1 2 Hamilton, W. D.; Axelrod, R.; Tanese, R. (1990). "Sexual reproduction as an adaptation to resist parasites (a review)". Proceedings of the National Academy of Sciences. 87 (9): 3566–3573. Bibcode:1990PNAS...87.3566H. doi: 10.1073/pnas.87.9.3566 . ISSN   0027-8424. PMC   53943 . PMID   2185476.
  9. Morran, L. T.; Schmidt, O. G.; Gelarden, I. A.; Parrish, R. C.; Lively, C. M. (2011). "Running with the Red Queen: Host-Parasite Coevolution Selects for Biparental Sex". Science. 333 (6039): 216–218. Bibcode:2011Sci...333..216M. doi:10.1126/science.1206360. ISSN   0036-8075. PMC   3402160 . PMID   21737739.